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Biomedical subjects

M A Nordstrom

Publications and source records attributed to M A Nordstrom.

At least 19 recordsLinked to original sources

Motor unit discharge and force tremor in skill- and strength-trained individuals.

We examined motor unit (MU) discharge properties (mean interspike interval, ISI, discharge variability, short-term synchronization, common drive) and force tremor in the first dorsal interosseous (FDI) muscle of five musicians (skill-trained), five weight-lifters (strength-trained) and six untrained subjects during low-force isometric abduction of the index finger. Mean MU ISI was slightly shorter in skill-trained subjects than in untrained subjects. Discharge variability of FDI MUs did not differ significantly between groups. The mean strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was different in skill-trained (0.22+/-0.02 s(-1), 162 MU pairs), untrained (0.32+/-0.02 s(-1), 199 MU pairs) and strength-trained subjects (0.44+/-0.03 s(-1), 183 MU pairs). FDI MU synchrony was weak and of equivalent strength in both hands of skill-trained subjects and the dominant (skilled) hand of untrained subjects. The stronger FDI MU synchrony in the non-dominant hand of untrained subjects was equivalent to that found in both hands of strength-trained subjects. The extent of common modulation of firing rates (common drive) was assessed for a subset of MU pairs and was weaker in skill-trained subjects (0.30+/-0.04, n=14) than untrained (0.43+/-0.3, n=14) and strength-trained (0.48+/-0.03, n=21) subjects. Force tremor was quantified for each hand in the same subjects during isometric index finger abduction at target forces of 0.5 N and 3.5 N. Tremor rms amplitude and peak power in the force frequency spectrum were significantly lower in skill-trained subjects than strength-trained subjects with the 3.5-N target force. The peak tremor frequency was similar in the three groups. The relatively more independent discharge of pairs of FDI MUs in skill-trained subjects was not responsible for the reduced tremor amplitudes in these subjects. Correlations between the overall extent of MU synchrony and common drive in FDI muscles and tremor measures obtained during the same experimental session were all non-significant. Differences in the central descending command signals are the most likely explanation for the more independent discharge of FDI MUs in skill-trained hands, while neural or peripheral muscular factors may be responsible for the weaker tremor.

Adolescent

Hemispheric differences in motor cortex excitability during a simple index finger abduction task in humans.

Transcranial magnetic (TMS) and electrical (TES) stimulation was used to assess the contribution of the corticospinal pathway to activation of the first dorsal interosseous muscle (FDI) in each hand of 16 right-handed subjects. TMS was applied at relaxed threshold intensity while the subject performed isometric index finger abduction at seven force levels [0.5 N to 50% maximal voluntary contraction (MVC)]. In a separate session, TES of equivalent intensity was applied to each hemisphere in 5 of these subjects while they performed the same force-matching protocol. In the resting state, mean threshold intensity for a muscle-evoked potential (MEP) in FDI using TMS was similar for the hemispheres controlling the dominant and nondominant hands. The size of the threshold MEPs in resting FDI after TMS and TES were also similar in each hand. With TMS, contraction-induced facilitation of the MEP in FDI was significantly larger when the nondominant hand was used for index finger abduction. In the pooled data, the nondominant/dominant ratio of MEP areas (normalized to the maximum M wave) ranged from 1. 7 in the weakest contraction (0.5 N) to 1.1 in the strongest (50% MVC). Eight subjects had significant differences between hands in favour of the nondominant hand, whereas in two subjects contraction-induced facilitation of MEPs was larger in the dominant hand. In five subjects for whom detailed motor unit data were available from a previous study, lateral differences in MEP facilitation were positively correlated with differences in FDI motor unit synchronization between hands. With TES, contraction-induced facilitation of the MEP was similar in each hand, suggesting that spinal excitability was equivalent on both sides. For the group of five subjects tested with both stimulation techniques, contraction-induced facilitation of the MEP was significantly larger after TMS than that obtained with TES when the contraction was performed with the nondominant hand, but not when the dominant hand was used to perform the task. We conclude that the extent of corticospinal neuron involvement in the command for simple index finger abduction in right-handed subjects is generally greater when the nondominant hand is used, compared with the same task performed with the dominant hand.

Adolescent

Relationship between motor unit short-term synchronization and common drive in human first dorsal interosseous muscle.

We assessed the strength of motor unit (MU) short-term synchronization and common fluctuations in mean firing rate (common drive) in the same pairs of MUs in order to evaluate whether these features of voluntary MU discharge arise from a common mechanism. Shared, branched-axon inputs, with the most important being widely divergent monosynaptic projections to motoneurons from motor cortical cells, are regarded as the principal determinants of MU short-term synchronization. It is not known to what extent these synaptic inputs are responsible for common drive behaviour of MUs. MU spike trains from 77 pairs of concurrently active MUs in first dorsal interosseous muscle of 17 subjects were discriminated with the high reliability needed for common drive analysis. For each MU pair, the data used for comparison of the two analyses of correlated MU discharge came from a single trial (1-5 min duration) of isometric abduction of the index finger. Linear regression revealed a weak, significant positive correlation between the strength of MU short-term synchronization and the strength of common drive in the MU pairs (r2 = 0.06, P < 0.05, n = 77), which was slightly stronger when MU pairs with broad synchronous peaks (> 20 ms) were excluded (r2 = 0.09, P < 0.05, n = 63). These data suggest that less than 10% of the variation in the strength of common drive exhibited by pairs of MUs could be accounted for by differences in the strength of MU short-term synchronization. These two phenomena are therefore likely to arise predominantly from separate mechanisms. At least under these task conditions, the widely divergent, branched-axon inputs from single corticospinal neurons which are important in the generation of MU short-term synchronization play only a minor role in the production of common drive of MU discharge rates.

Adolescent

Responses of human masseter motor units to stretch.

1. The reflex responses to stretch were studied in single motor units and the surface electromyogram in human masseter. 2. Controlled stretches of the isometrically contracting jaw-closing muscles evoked short-latency (10-15 ms) and long-latency (35-70 ms) excitatory reflex responses in the masseter surface electromyogram. 3. The majority (65%) of tonically active masseter motor units were excited in both short- and long-latency phases of the reflex. The timing of the stimulus determined whether the unit discharged in the short- or long-latency phase. If a non-tonically active motor unit was recruited by the stimulus, it invariably discharged in the long-latency phase. 4. Although short-latency responses were strongly time-locked to the stimulus, there was very little shortening of interspike intervals (ISIs) in this phase of the reflex. The shortening of ISIs was more prominent and prolonged during the long-latency phase, which explains why this phase produces most of the reflex force changes following the stretch. 5. Within pairs of concurrently active motor units there was a tenfold range in the size of the short-latency response to the same stretch. 6. A substantial proportion (35%) of the twenty-two masseter motor units tested had no statistically significant short-latency reflex response. 7. In contrast to other human muscles, there was no functional connection between a population of Ia afferents and some masseter motoneurons. There are two possible explanations for this result. The short-latency, presumably monosynaptic, Ia afferent inputs may not be uniformly distributed to human masseter motoneurons. Alternatively, these inputs may be subject to tonic presynaptic inhibition that is not uniformly distributed throughout the masseter motoneuron pool.

Adult

Fatigue of jaw muscles and speech mechanisms.

Histochemical studies show that the distribution of fiber types in human jaw muscles is different from that in various limb muscles, no doubt representing different functional demands as well as a different embryological derivation. Jaw-closing muscles appear more resistant to fatigue than limb muscles with intermittent maximal contractions. Endurance of continuous isometric biting is limited by pain. Masseter motor unit fatigability in sub-maximal contractions is similar to the limb muscles. There are few physiological data for the jaw-opening muscles. The distribution of fiber types in human speech muscles is consistent with the high speeds of contraction that must be used in phonation. Although clinical syndromes of fatigue of speech muscles are recognized, there is little direct information on the fatigability of the muscle fibers themselves.

Animals

Limitations of the surface electromyography technique for estimating motor unit synchronization.

Motor unit synchronization was estimated from the surface electromyograms (EMG) of the first dorsal interosseus muscle of human volunteers by a simplified surface-EMG technique (Milner-Brown et al. 1973, 1975). Single motor units were identified from intramuscular recordings and were used to obtain a spike-triggered average of the surface-EMG. The discharge rate of a reference motor unit was controlled at two levels (high and low), and the effect of motor unit activity on the surface-EMG estimate of synchronization was studied in 56 motor units. The surface-EMG estimate of motor unit synchronization was significantly higher when the reference motor unit discharged at the high rate than when it discharged at the low rate. A regression analysis indicated that the synchronization ratio calculated from the surface EMG was significantly correlated with the level of EMG activity in the muscle. Motor unit synchronization was also estimated from surface-EMG measurements that were derived by computer simulation. The simulation permitted manipulation of motor unit activity (discharge rate and recruitment) with a complete absence of synchrony among the units in the pool. The stimulated surface-EMG index was influenced by an artifact associated with signal rectification, and this effect changed non-monotonically with motor unit activity. Furthermore, the increase in the motor unit activity reduced the signal-to-noise ratio of the spike-triggered surface EMG average, and consequently decreased the sensitivity of the surface-EMG index as an estimate of motor unit synchronization. We conclude that the simplified surface-EMG method (Milner-Brown et al. 1973, 1975) does not provide a useful index of motor unit synchronization due to its inability to accurately distinguish the synchronization from methodological effects related to a rectification artifact and variation in the signal-to-noise ratio.

Computer Simulation

Discharge patterns of tonically firing human motoneurones.

We have attempted to reconcile the different patterns of distribution of interspike intervals that are found in motoneurones made to discharge by intracellular injection of constant current in reduced animal preparations and by voluntary control in human subjects. We recorded long spike trains from single motor units in three human muscles made to discharge at constant mean frequencies with the help of auditory and visual feedback. The distribution of interspike intervals in each spike train was analysed quantitatively. We found that the different pattern of discharge of the human motor units could be accounted for when due allowance was made for the variability of the drive to the human motoneurone which arose because of the feedback process used to maintain the target frequency. A model testing this hypothesis gave results that were qualitatively consistent with the human data.

Acoustic Stimulation

Influence of handedness on motor unit discharge properties and force tremor.

Discharge properties of motor units (MUs) in the first dorsal interosseous muscle (FDI) were studied in the dominant and non-dominant hands of six right-handed (RH) and six left-handed (LH) individuals. MU discharge rates and variability were similar in each hand in RH (186 MUs) and LH (160 MUs) subjects. MU synchronization was less prominent in the dominant hand of RH subjects, with 51% (45/88) of cross-correlograms of MU discharge having significant central peaks, compared with 81% (90/111) for the non-dominant hand. The strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was weaker in the dominant hand of right-handers (0.23 +/- 0.03 s-1 vs 0.39 +/- 0.03 s-1), and synchronous peaks from that hand were slightly broader. Four of six RH subjects had significant differences in synchronization between hands (weaker in dominant hand). In contrast, left-handers had similar incidence (80 vs 82%, n = 161) and strength (0.41 +/- 0.03 s-1 vs 0.37 +/- 0.03 s-1) of MU synchrony in dominant and non-dominant hands. No LH subject had a significant difference in synchronization between hands. Force tremor was quantified in each hand in the same subjects during isometric abduction of FDI at 0.5 N and 3.5 N, and directly correlated with the extent of MU synchronization in the muscle. Tremor root mean square amplitude was similar in dominant and non-dominant hands. Power spectral analysis of the tremor force revealed that the peak frequency in the power spectrum was not influenced by handedness, but power at the peak frequency was higher in the non-dominant hand of RH subjects. Correlations between MU discharge variability and synchrony with measures of tremor amplitude were weak. The reduced MU synchronization in the dominant hand of right-handers may reflect a more restricted distribution of direct projections from motor cortical neurons within the FDI motoneuron pool, or reduced excitability of the cortical neurons during the task. These differences in MU synchronization, however, had an insignificant influence on the magnitude of physiological tremor in the FDI.

Adult

Spike-train acquisition, analysis and real-time experimental control using a graphical programming language (LabView).

A solution is described for the acquisition on a personal computer of standard pulses derived from neuronal discharge, measurement of neuronal discharge times, real-time control of stimulus delivery based on specified inter-pulse interval conditions in the neuronal spike train, and on-line display and analysis of the experimental data. The hardware consisted of an Apple Macintosh IIci computer and a plug-in card (National Instruments NB-MIO16) that supports A/D, D/A, digital I/O and timer functions. The software was written in the object-oriented graphical programming language LabView. Essential elements of the source code of the LabView program are presented and explained. The use of the system is demonstrated in an experiment in which the reflex responses to muscle stretch are assessed for a single motor unit in the human masseter muscle.

Action Potentials

Reduced motor unit activation of muscle spindles and tendon organs in the immobilized cat hindlimb.

Six weeks of limb immobilization of a healthy muscle (cat tibialis posterior) at a short length resulted in a significant reduction of mean fiber area for all fiber types (I, 71% of control; IIa, 77% of control; IIb, 79% of control), whereas fiber type proportions were unchanged. For motor units, there was a reduction in peak tetanic force (type slow > fast fatigue resistant > fast fatigable); an increase in the twitch-to-tetanus ratio for fast fatigue-resistant and slow units; and no effect on the twitch force, twitch time course, or fatigability. The reduction in peak force was greater than expected because of fiber atrophy in slow units. Immobilization had a minimal effect on muscle spindle afferent (Ia and spindle group II) responses to a ramp-and-hold stretch of the passive muscle. Tendon organ (Ib) afferents had an increased responsiveness to stretch after immobilization but only when the muscle was stretched from a short resting length. However, immobilization reduced the modulation of muscle afferent discharge in response to tetanic contractions of single motor units. The decline in responsiveness of spindles was a result of the reduced tetanic force of motor units. In contrast, tendon organs in immobilized muscle were twice as likely to convey no information on the contraction of a single motor unit and were more likely to be unloaded, suggesting that immobilization caused the functional denervation of some muscle fibers. Thus the responses of muscle spindles and tendon organs in immobilized muscle reflected atrophic changes in extrafusal fibers but did not provide evidence for substantial disturbance of receptor function.

Animals

A new approach to the estimation of post-synaptic potentials in human motoneurones.

A new method is described for estimating the shape of the compound post-synaptic potentials evoked by stimuli in human motoneurones. The method is based on changes in the duration of the interspike intervals in motor-unit spike trains that are time-locked to the stimulus. This is particularly helpful in estimating the profile of long-latency slow rise-time post-synaptic potentials that are difficult to estimate with other methods. The method is simple to apply and the analysis is readily implemented on a personal computer.

Electric Stimulation

Adaptation of cat motoneurons to sustained and intermittent extracellular activation.

1. The main purpose of this study was to quantify the adaptation of spinal motoneurons to sustained and intermittent activation, using an extracellular route of stimulating current application to single test cells, in contrast to an intracellular route, as has been used previously. In addition, associations were tested between firing rate properties of the tested cells and other type (size)-related properties of these cells and their motor units. 2. Motoneurons supplying the medial gastrocnemius muscle of the deeply anaesthetized cat were stimulated for 240 s with microelectrodes which passed sustained extracellular current at 1.25 times the threshold for repetitive firing. Many cells were also tested following a rest period with intermittent 1 s current pulses (duration 600 ms) at the same relative stimulus strength. Cell discharge was assessed from the EMG of the motor unit innervated by the test neuron. The motoneurons and their motor units were assigned to four categories (i.e. types FF, FR, S and F; where F = FF + FR) based on conventional criteria. In all, twenty F (16 FF, 4 FR) and fourteen S cells were studied with sustained stimulation. Thirty of these cells (17 F, 13 S) and an additional two cells (1 F, 1 S) were studied with intermittent stimulation. 3. The mean threshold current required for sustained firing for a period of > or = 2 s was not significantly different for F and S cells. However, most of the other measured parameters of motoneuron firing differed significantly for these two cell groups. For example, at 1.25 times the threshold current for repetitive firing, the mean firing duration in response to 240 s of sustained activation was 123 +/- 88 s (+/- S.D.) for F cells vs. 233 +/- 19 s for S cells. These values were significantly longer than those from a comparable, previously reported study that employed intracellular stimulation. With intermittent stimulation, the firing durations of F and S cells were not significantly different from each other. 4. All cells exhibited a delay from the onset of current to the first spike, followed by a brief accelerating discharge that was followed by a slower drop in firing rate. Some cells (21 of 34 with sustained activation; 20 of 32 with intermittent) exhibited doublet discharges (interspike intervals < or = 10 ms) that were intermingled with the more predominant singlet discharges. Doublets were more common in the S cell type.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

Estimating the strength of common input to human motoneurons from the cross-correlogram.

1. The relationship between the motor unit discharge pattern (rate and variability) and synchronization of motor unit pairs was studied in the first dorsal interosseus muscle of human subjects. In separate trials of up to 4 min duration, subjects voluntarily controlled the mean discharge rate of an identified motor unit at one of several prescribed rates (range 7.5-17.5 Hz). 2. The effect of discharge rate on the synchronous peak in the cross-correlogram was examined in eighty motor unit pairs from six subjects. Five commonly used synchronization indices were used to quantify synchrony in the cross-correlograms constructed from different discharge-rate trials. For each synchronization index, the apparent magnitude of synchrony increased at lower motor unit discharge rates. The synchronization indices were not equally sensitive to discharge rate; increases in the different indices ranged from 72 to 494% between the highest and lowest discharge rates. 3. A model of the membrane potential trajectory underlying rhythmic motoneuron discharge was used to explain the observed increase in the magnitude of the synchronization indices at lower discharge rates. The essential feature of this model is that the probability of a common-input EPSP causing a synchronous discharge in two motoneurons is independent of discharge rate. This means that the number of synchronous action potentials in excess of chance in any trial depends on the properties of the common-input EPSPs and the duration of the trial, but is not related to motor unit discharge rates. The model also demonstrated that when the excess synchronous counts are normalized to motor unit discharge rate, or baseline counts in the histogram (as in the conventional synchronization indices), the magnitude of the index increases when the motor unit discharge rates are low. 4. The strength of common input to motoneurons could be misinterpreted if conventional synchronization indices are used because of discharge-rate effects. The model was used to derive an index of the strength of common input to motoneurons (CIS) that was independent of motor unit discharge rate. CIS is the frequency of synchronous action potentials in the motor unit pair in excess of those expected due to chance (calculated during periods of tonic discharge in both units). The mean CIS in first dorsal interosseus motor unit pairs ranged from 0.052 to 1.005 extra synchronous action potentials per second across subjects. 5. Discharge variability was correlated with each of the synchronization indices and the CIS.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

Instability of motor unit firing rates during prolonged isometric contractions in human masseter.

The firing patterns of up to 4 concurrently active masseter motor units were studied with intramuscular electrodes during a continuous isometric contraction of 15 min duration, in which the subject maintained the mean firing rate of one selected unit at 10 Hz. With this paradigm the net excitation (i.e. mean firing rate) of one unit in the muscle was controlled. This served as the reference for the functional state of other active units during the prolonged contraction. With the mean firing rate of one unit in the muscle fixed, 58% of other active units showed a slow, statistically-significant change in mean firing rate over the 15 min. The initial firing rate of the units did not influence the change in rate. The original firing rate hierarchy, which in short-term contractions reflects the recruitment order, was altered during the prolonged contraction. The explanation for these differential changes in motoneuron net excitation is not clear; they could be intrinsic to the motoneurons or perhaps mediated by reflex pathways. The selective facilitation or suppression of some motor units with continuous activation means that the original size-structured combination of motor units can be modified during a prolonged contraction.

Adolescent

Discharge variability and physiological properties of human masseter motor units.

Intramuscular electrodes were used to study discharge variability in motor units of human masseter whose physiological properties were determined using spike-triggered averaging. Subjects voluntarily controlled the mean firing rate of a selected motor unit at 10 Hz for 15 min of continuous activation. Discharge variability was assessed at the beginning and end of this period. In 81% of units, the discharge variability at a mean interspike interval (ISI) of 100 ms increased after 15 min of continuous activity. There was a wide range of discharge variability within the population of masseter units studied, but no significant correlations were found between initial discharge variability and recruitment threshold, twitch tension or time-to-peak tension (TTP). There was, however, a significant correlation between motor unit fatigability and its initial discharge variability. This represents a link between the motoneuron and the functional properties of the muscle fibers in innervates.

Adolescent

Synchronization of motor units in human masseter during a prolonged isometric contraction.

1. The action potentials of concurrently active motor units in human masseter were recorded on several fine-wire intramuscular electrodes and synchronization was assessed by cross-correlation of their firing times. 2. Long recording sessions and a test of significance for peaks in histograms were used to enhance detection of synchrony. 3. Synchrony in masseter units was found to be very weak, and was observed consistently only in histograms with a large number of counts. However, synchrony was widespread within the motoneurone pool; of the unit pairs whose cross-correlation histograms contained more than 100 counts per 1 ms bin, 89% were significantly synchronized. 4. The strength of synchrony between pairs of units did not change consistently during 15 min of continuous activity. 5. It was concluded that most, if not all, low-threshold masseter motoneurones have some common synaptic inputs, although the origin of those inputs remains unclear.

Action Potentials

Fatigue of single motor units in human masseter.

The spike-triggered averaging technique was used to determine the time course and extent of fatigue of single motor unit twitches in the human masseter. This is the first report of a fatigue test having been applied to masseter motor units in either animals or humans. The human masseter was found to be comprised predominantly of fast-twitch motor units with a broad spectrum of fatigability. Very few physiological type S units were found, despite histochemical evidence for a substantial population of type I fibers in the masseter. In addition, there was no significant correlation between fatigability and either twitch amplitude or contractile speed in the motor units studied. The latter observations are consistent with the unusual histological features of the masseter. Comparison with other human fatigue data suggests that the extent of fatigue in the present population of masseter motor units after approximately 3,000 activations is similar to that reported for populations of units in first dorsal interosseous and medial gastrocnemius.

Adolescent

Effect of motor unit firing pattern on twitches obtained by spike-triggered averaging.

The spike-triggered averaging technique (STA) was used to determine the twitch profile of single motor units in human masseter during a voluntary isometric contraction. The effect of the immediate firing pattern of the unit on the twitch was assessed with a computer program which scanned the unit discharge records and selected valid trigger spikes for the averager on the basis of the interspike intervals preceding and following the trigger spike. Successive averages from the same data using different interval parameters revealed progressively more fusion of twitches as the instantaneous firing rate increased. When the data were averaged with interval parameters similar to those used in earlier studies, some fusion of the twitch profile was also evident. It is therefore likely that the degree of fusion of human motor unit twitches obtained by STA has been underestimated in the past. It is further concluded that masseter motor units are sufficiently fast-contracting to allow a relatively unfused twitch profile to be obtained with STA, provided trigger spikes are subjected to appropriate rate control.

Adolescent